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Trachylepis gonwouoi Allen, Tapondjou, Welton & Bauer sp. nov. Holotype. California Academy of Sciences, San Francisco, CA: CAS 261092, adult male, collected near the town of Mudeka near Mt. Cameroon, Southwest Region, Cameroon, 4.125��N, 9.491��E, 16 June 2014, by Kaitlin Allen, Walter Tapondjou and Luke Welton (Fig. 2). Paratypes. Monte L. Bean Life Science Museum, Provo, Utah and California Academy of Sciences, San Francisco, CA: CAS 261090, juvenile, town of Buea near Mt. Cameroon, Southwest Region, Cameroon, 4.160��N, 9.228��E; CAS 261091, female and BYU 573522, female, town of Idenau, Southwest Region, Cameroon 4.183��N, 9.000��E; CAS 261093, juvenile, BYU 573527, juvenile, BYU 573861, juvenile, BYU 573529, female, and BYU 573530, male, town of Nyasoso on Mt. Kupe, Southwest Region, Cameroon, 4.823��N, 9.670��E; 14 June 2016 ��� 2 July 2016; coll. Kaitlin Allen, Walter Tapondjou and Luke Welton. (Fig. 3 and Fig. 4) Other material referred. Additional specimens referable to the new species based on mitochondrial genotype are: United States National Museum of Natural History (USNM) 584340, town of Impongui, Likouala Province, Republic of the Congo, 1.079��N, 17.036��E; Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK) 68962 and 75386, Mt. Nlonako, Littoral Region, Cameroon, 4.906��N, 9.951��E; and ZFMK 87654, Campo Reserve, South Province, Cameroon, 2.337��N, 9.857��E; Museum f��r Naturkunde [no voucher collected], Mt. Nlonako, Littoral Region, Cameroon, 4.906��N, 9.951��E; and [no voucher collected], Rumpi Hills, Southwest Region, Cameroon, 4.898��N, 9.119��E (Fig. 4). Diagnosis. Trachylepis gonwouoi sp. nov. can be distinguished from other Trachylepis in West and Central Africa by the following characteristics, number of keels on dorsal scales (3���5); moderate SVL (maximum size of 80 mm); number of scale rows at midbody (28���34); number of supracilliaries (6���10); presence of a well defined lateral white stripe extending from beneath eye to insertion of hind limb, bordered by black; and ventral color in life bright blue-green. Description of the holotype. Adult male specimen; tail regenerated; right parietal scale missing, replaced by scar tissue; some damage to scales on right side of neck; SVL 71 mm; regenerated tail length 101 mm, approximately 1.4 times SVL; head depth 7.6 mm; orbit width 5.7 mm; eye height 3.4 mm; distance from anterior portion of tympanum to tip of snout 16 mm; distance from rictus to tip of snout 13.1 mm; supralabials 7 (L) 8 (R), fifth supralabial located under eye, widened; infralabials 8 (L) 9 (R); 4 supraoculars; supracilliaries 7 (L) 6 (R); 2 supranasals; 2 loreals, anterior roughly half the size of posterior; prefrontals contact at a medial point; supranasals narrowly separated by rostral and frontonasal; nostril bisects nasal scale into postnasal and prenasal; nuchal scales in broad contact; frontoparietals divided by interparietal plate; frontoparietals of equal length; parietals in broad contact; tympanic aperture large and vertically elongated with two well-developed anterior lobules on each side (Fig. 5); right forelimb length 26 mm; right hind-limb length 34.2 mm (measured to the terminus of the longest digit); axilla���groin distance 37.1 mm; limb scales keeled; digits I and IV entirely missing on left hand; subdigital lamellae Toe IV 15 (L) 18 (R); subdigital lamellae Finger IV 10 (R); scales on ventral surface of manus and pes smooth; 30 transverse scale rows at midbody; dorsal scales with 3���5 keels; 49 paravertebral scales between nuchals and base of tail; 54 ventral scales between the first pair of postmentals and cloaca; 6 precloacal scales smooth, medioventrally widened; caudal scales medio-ventrally widened, smooth; regenerated caudal scales large, carination less distinct. Coloration (preserved specimen): body dorsum uniformly brown with a lateral white stripe from under the eye to the insertion of the hind limb. White stripe 1 mm (one scale row) wide at midbody, widening to 1.5 mm (1.5 scale rows) near forearm insertion, bordered by thin black stripes, about 0.5 mm thick. Ventrally uniform gray-blue from snout to tail tip, limbs ventrally darker gray-blue, ventral surface of manus and pes black (Fig. 2). Color in life: Dorsal surface solid brown with no mottling; flanks red-brown to middle of flank; white lateral line (1 to 1.5 mm thick) bordered by two thinner black lines (0.5 mm thick) on flank from snout to insertion of hind limb; ventral quarter of flank and venter from tip of snout to tip of tail blue-green; ventral surface of manus and pes black; eyelid borders yellow and contrasting with surrounding brown scales; eyelids black; eye color dark brown; limbs red-brown; feet darker brown than legs or dorsum (Fig. 6). Variation. Variation among members of the type series of Trachylepis gonwouoi sp. nov. in mensural and meristic characters are reported in Table 2. The range of variation for each character is reported here: SVL of adults, 71���80 mm; ratio of tail length to body length (for original tails only), 1.787���1.90; ratio of head length to body length, 0.158���0.275; number of scale rows at midbody, 28���34; number of paravertebral scales from nuchals to base of tail, 47���53; number of ventrals from first post-mental pair to the anal plates, 51���61; number of subdigital lamellae for Toe IV, 10���19; number of subdigital lamellae for Finger IV, 9���16; number of supralabials, 6���8; maximum number of keels on dorsal scales, 3���5; supranasials contact at a point or have no contact; prefrontals contact at a point or have no contact; number of supracilliaries, 6���10. Etymology. This species is a patronym honoring Dr. LeGrand Gonwouo Nono, the first native-born Cameroonian herpetologist, in recognition of his work to increase the knowledge of herpetology and conservation of endangered reptiles and amphibians throughout Cameroon. Vernacular name. Gonwouo���s skink (English); Scinque de Gonwouo (French). Distribution. We collected Trachylepis gonwouoi sp. nov. from two areas in the Southwest Province of Cameroon, on Mt. Cameroon, near the towns of Buea (4.160��N, 9.228��E), Idenau (4.183��N, 9.000��E) and Mudeka (4.125��N 9.491��E) and on Mt. Kupe near the town of Nyasoso (4.823��N, 9.670��E). Specimens from the United States National Museum of Natural History (USNM) and the Natural History Museum of Geneva (MHNG) were collected from Mt. Nlonako (4.905��N, 9.951��E), Campo (2.337��N, 9.857��E), and Rumpi Hills (4.898��N, 9.119��E). One specimen has also been collected in the Likouala province of the Congo near the town of Impongui (1.079��N, 17.306��E). The total distance between the two most distant localities is 1008 kilometers (Fig. 4). Natural history. Trachylepis gonwouoi sp. nov. was associated with disturbed habitat in agricultural and residential areas as well as secondary forest. It was often found in the leaf litter or on the trunks of cocoa trees. Specimens from Cameroon were found at elevations from 50 to 1050m. This species is syntopic with T. affinis and T. maculilabris., Published as part of Allen, Kaitlin E., Tapondjou, Walter P., Welton, Luke J. & Bauer, Aaron M., 2017, A new species of Trachylepis (Squamata: Scincidae) from Central Africa and a key to the Trachylepis of West and Central Africa, pp. 255-269 in Zootaxa 4268 (2) on pages 258-261, DOI: 10.11646/zootaxa.4268.2.5, http://zenodo.org/record/580227

Varanus samarensis Koch et al. 2010 Synonymy. Varanus cumingi samarensis Koch et al., 2010. Zootaxa 2446, 19 ��� 23. Holotype.��� ZFMK 64713, subadult male, collected by M. Gaulke 1989, San Augustin near Gandara, Samar Island, Philippines. New material examined.���Samar (1 specimen): KU 310870 (CDS Field No. 2654), adult male, collected by Cameron D. Siler, 13 October 2007, Barangay San Rafael, Municipality of Taft, Eastern Samar Province, Samar Island, Philippines; Bohol (1 specimen): KU 335263 (CDS Field No. 4768), adult male, collected by Cameron D. Siler, 0 4 August 2009, Barangay Danicop, Municipality of Sierra Bullones, Bohol Province, Bohol Island, Philippines. Other material. ���Refer to Koch et al. (2010 b). Diagnosis. ���Following the findings of Koch et al. (2010 b), Varanus samarensis can be distinguished from other members of the V. salvator Complex by a combination of (1) distinctive dorsal color pattern of 5���8 transverse bands of yellow ocelli or spots through the trunk; (2) a dark (black) head dorsally, with symmetrical yellow markings; (3) a variable black stripe in the temporal region, occasionally bordered by a bright (yellow-white) stripe ventrally; (4) the absence of a medio-dorsal stripe on the lateral surfaces of the nuchal region and trunk; (5) narial openings positioned approximately 2.5 times closer to the tip of the snout than to the ocular. Comment and justification. ���Although within the same PAIC (Brown & Diesmos 2009), the Samar���Leyte versus Mindanao Island biogeographical boundary (informally referred to as the ���Dinagat���Caraga Filter Zone���) has been evoked in several previous studies to explain the existence of deep phylogenetic splits between the northern and southern reaches of the Mindanao faunal region (Steppan et al. 2003; Hosner et al. 2013, 2014; Brown et al. 2013, 2014). We therefore do not find it surprising that Samar, Leyte, Bohol (and, presumably, smaller associated lanmasses) have a distinctive species of monitor lizard that apparently is isolated from Mindanao���s Varanus cumingi (Welton et al. 2013 a, b). Distribution. ���See Koch et al. (2010 b) for additional data about variation, ecology, and distribution., Published as part of Welton, Luke J., Travers, Scott L., Siler, Cameron D. & Brown, Rafe M., 2014, Integrative taxonomy and phylogeny-based species delimitation of Philippine water monitor lizards (Varanus salvator Complex) with descriptions of two new cryptic species, pp. 201-227 in Zootaxa 3881 (3) on page 221, DOI: 10.11646/zootaxa.3881.3.1, http://zenodo.org/record/226203, {"references":["Koch, A., Gaulke, M. & Bohme, W. (2010 b) Uravelling the underestimated diversity of Philippine water monitor lizards (Squamata: Varanus salvator complex), with the description of two new species and a new subspecies. Zootaxa, 2446, 1 - 54.","Brown, R. M. & Diesmos, A. C. (2009) Philippines, Biology, In: Gillespie, R. & Clague, D. (Eds.), Encyclopedia of Islands. University of California Press, Berkeley, pp. 723 - 732.","Steppan S. J., Zawadzki C, Heaney L. R. (2003) Molecular phylogeny of the endemic Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic archipelago. Biological Journal of the Linnaean Society, 80, 699 - 715.","Welton, L. J., Siler, C. D., Linkem, C. W., Diesmos, A. C., Diesmos, M. L., Sy, E. & Brown, R. M. (2013 a) Dragons in our midst: Phyloforensics of illegally traded Southeast Asian monitor lizards. Biological Conservation, 159, 7 - 15.","Welton, L. J., Siler, C. D., Oaks, J. R., Diesmos, A. C. & Brown, R. M. (2013 b) Mulilocus Phlogeny and Bayesian estimates of species boundaries reveal hidden evolutionary relationships and cryptic diversity in Southeast Asian monitor lizards. Molecular Ecology, 22 (13), 3495 - 3510."]}

Varanus bangonorum sp. nov. Figs. 3, 6���9 Holotype. PNM 9798 (formerly University of Kansas Natural Biodiversity Institute [KU] 335744; Field no. RMB 17757), sub-adult male, collected by SLT and RMB, 16 July 2013, 230 m above sea level (12.787806, 120.915972; WGS- 84), Sitio Aruyan, Barangay Malisbong, Municipality of Sablayan, Occidental Mindoro Province, Mindoro Island, Philippines. Paratopotypes. KU 335742 (RMB Field No. 17720), juvenile, collected 11 July 2013. Paratypes. KU 335743 (RMB Field No. 17824), juvenile, Lake Libuao, Barangay Malisbong, Municipality of Sablayan, Occidental Mindoro Province, Mindoro Island, Philippines; PNM 9799 (formerly KU 308403), 308437 (ELR Field Nos. 843, 877), juvenile and adult female, Sitio Ulasan, Barangay Harrison, Municipality of Paluan, Occidental Mindoro Province, Mindoro Island, Philippines; KU 305151, 305161, 305163 (CDS Field Nos. 714, 715, 648), juveniles, Barangay Tinogboc, Municipality of Caluya, Antique Province, Semirara Island, Philippines. Other Material. See Specimens Examined Section Etymology. The specific epithet, bangonorum, is a plural noun derived from the name of the indigenous Bangon peoples of Mindoro Island, who reside along the rivers and central Cordillera mountain system in Oriental Mindoro Province. Anthropologists have shown the Bangon tribal group to be culturally distinct from other tribal groups inhabiting Mindoro (Tweddell 1970; Mangyan Heritage Center). Suggested common name: Bangon Monitor Lizard. Diagnosis. Varanus bangonorum can be distinguished from congeners by (1) distinct, relatively large dark spots or blotches in the gular region; (2) low counts of dorsal gular scales; (3) low counts of dorsal trunk scales; and (4) low counts of scales around the neck anterior to the gular fold. Additionally, this distinct lineage is biogeographically circumscribed on the deepwater islands of Mindoro and Semirara. Comparisons. This new species in phenotypically most similar to Varanus marmoratus, from which it can be distinguished by the presence of distinct, dark and irregularly-shaped spots or blotches in the gular region (versus speckling and transverse bands; Fig. 3), and an allopatric distribution on the islands of Mindoro and Semirara (versus Luzon and Lubang islands, and the Batanes and Babuyan island groups; Figs. 1, 2). Additionally, although the new species is phenotypically similar to V. marmoratus, it is not closely related to this species (Fig. 2, phylogeny). Varanus bangonorum can be distinguished from its closest but allopatric relative, V. palawanensis, by having generally fewer rictus���rictus scales (mean = 56 �� 4, versus 62 �� 4), fewer dorsal scales from the tympanum to the gular fold (mean = 26 �� 2, versus 32 �� 3), fewer dorsal scales from the nuchal fold to the hind limb insertion (mean = 80 �� 9, versus 96 �� 3), fewer total dorsal scales (mean = 106 �� 10, versus 128 �� 3), fewer scales around the neck anterior to the gular fold (mean = 76 �� 6, versus 87 �� 4), and by having distinct, dark irregular spots or blotches in the gular region (versus irregular speckling with anterior transverse bands). Of the remaining geographically proximate species of the V. salvator Complex, this new species can be distinguished from V. nuchalis by a distribution on Mindoro and Semirara islands (versus Visayan islands), generally fewer midbody scales (mean = 136 �� 9, versus 150 �� 7), generally more dorsal scales from the tympanum to the gular fold (mean = 26 �� 2, versus 23 �� 2), and the presence of distinct dark spots or blotches in the gular region (versus a uniform dark gular coloration). Additionally, this new species can be distinguished from the allopatric Bicol species V. dalubhasa by having distinct, irregularly shaped spots or blotches (versus variable speckling and faint anterior transverse bands) in the gular region. Lastly, V. bangonorum exhibits entirely unique haplotypes or haplotype networks, relative to all other members of the V. salvator Complex (see Welton et al. 2010 a, b). We have constrained our morphological analyses to the geographically most proximate taxa due to previous studies (Koch et al. 2007, 2010 b) demonstrating their distinctiveness relative to the remaining diversity within the V. salvator Complex. Description of holotype. A sub-adult male, hemipenes not everted; snout���vent length 343 mm; tail 601 mm; head relatively slender, length 65.9 mm, width 38.2 mm, and depth at eye 28.5 mm; snout length 34.0 mm, 51.6% head length; snout rounded anteriorly; narial openings 4.3 mm, ovular, posteriorly elevated, encircled by 9 L/ 9 R small polygonal scales; snout elevated sharply just anterior to narial region, then continuing gradually to ocular region; narial region only moderately elevated above horizon of snout; cranial table squarish, slightly wider than long. Head scales ovular to polygonal (Fig. 9), heterogenous in size with the smallest occuring on lateral margins of cranial table and directly surrounding pinneal scale, and largest occuring dorso-medially from tip of snout to pineal region; supraoculars 7 L/ 6 R, subrectangular and elongated transversely; supralabials 52, slightly larger than head scales; infralabials 52, smaller than supralabials; both supra- and infralabials increasing in size anteriorly; nuchals large, circular, generally homogenous in size throughout nuchal region except for directly posterior to cranial table; lateral nuchals small and granular, approximately �� the size of dorsals; sutures between scales larger than those on head; nuchals 25, in semi-regular transverse rows from posterior margin of tympanum to forelimb insertion; dorsal trunk scales arranged in 77 semi-regular transverse rows from fore- to hindlimb insertions, ovular, generally smaller than nuchals, and decreasing in size posteriorly to hindlimb insertion as well as laterally, with smallest scales occurring adjacent to limb insertions; midbody scales 131; axilla-groin distance 157 mm; dorsal limb scales ovular, decreasing in size distally and ventrally; fore- and hindlimbs 55 and 82 mm, respectively, 16.0% and 23.9% snout���vent length, respectively, 24.2 and 36.8 mm diameter at insertions, respectively; limb scales present in semiregular rows; scales of manus and pes smaller and more irregular in shape, ranging from circular to nearly square dorsally; supradigitals rectangular dorsally, ovular laterally, with terminal scale nearly twice as large as others; digits terminate in long, slender recurved claws; caudals ovular, small and granular anteriorly, becoming rectangular posteriorly; dorsal keel composed of paired, posteriorly elevated scales commenses 37.7 mm from hindlimb insertion, continuing to tail terminus with scales gradually decreasing in size; caudal scales moderately keeled, in regular transverse and longitudinal rows laterally, decreasing in size posteriorly. Ventrals less variable than dorsals; gular scales generally rectangular anteriorly, bcoming more ovular towards gular fold; smallest scales medially in anterior third of gular region, with largest scales just anterior to gular fold but followed posteriorly by three transverse rows of smaller, more granular gulars; scales around the neck anterior to the gular fold 67, mid-gular 76; gular scales in 70 irregular rows from tip of snout to gular fold; ventrals of fore- and hindlimbs circular to polygonal, with those of forelimbs slightly smaller than those of hindlimbs; precloacals similar to those of hindlimbs; ventral trunk scales from gular fold to hind limb insertion 83, ovular anteriorly, becoming rectangular posteriorly, present in semi-regular rows; scales decreasing in size laterally and at hindlimb insertions; ventral caudal scales rectangular, smallest just posterior to cloacal opening; scale surface slightly rounded anteriorly, becoming sharply keeled and more longitudinally elongate posteriorly. Measurements of holotype (in mm). SVL 343; tail length 601; head length 65.9, width 38.2, depth at eye 28.5; eye���naris distance 22.2; naris���snout distance 11.6; rictus���rictus scales 52; scales around tail base 91; scales around tail 1 / 3 distance from base 47; mid-body scales 131; gular scales 70; ventrals from gular fold to hind limb insertion 83; total ventral scales 153; dorsals from tympanum to gular fold 25; dorsals from gular fold to hind limb insertion 77; total dorsals 102; supralabials 52; scales around the neck anterior to gular fold 67; and enlarged supraoculars 7 L/ 6 R. Coloration of holotype in preservative. Dorsal surfaces of body, head and limbs black; head nearly uniform black, with dark brown accents dorsal to narial openings and with white pineal scale; trunk with five transverse rows of white blotches; tail with nine irregular bands comprised of diffuse white speckling; forelimbs with sparse white speckling, becoming more regular ventrally; hindlimbs more speckled than forelimbs, and with more speckling ventrally; dorsal surfaces of manus and pes with few white speckles, but surface of pes with slightly more; terminal scale on digits with conspicuous white spot. Lateral coloration of head consisting of vague, anterior dark brown band extending ventrally from dark brown dorsal markings to narial openings; post-orbitally, lateral surface with longitudinal white stripe extending from the posterior margin of ocular to tympanum (Fig. 9); lateral coloration of the nuchal region consists of bright white mottling, gradually increasing in dorsal extent posteriorly, and terminating at forelimb insertion; lateral portions of trunk with variable white blotches and speckling, with greatest concentration directly posterior to forelimbs, the distribution of which does not correspond to dorsal banding; terminal half of tail solid black dorsally. Ventral coloration predominately yellow-white throughout; gular region with characteristic dark blotches, becoming more prominent posteriorly; anterior portions of gular region with three triangular projections extending medially; scapular region with irregular dark blotches; trunk with irregular dark triangular projections extending medially from margins, and generally not connecting with projections from opposite side; forelimbs and hindlimbs almost uniform white at insertions, but with increasing dark mottling laterally and in distal portions; manus and pes dark gray; venter of tail similar to that of trunk, but with triangular projections connecting and forming vague bands posteriorly; terminal third of tail solid black ventrally. Variation. Our small type series exhibits some color variation but specimens are generally phenotypically similar to one another. Four specimens from Mindoro, including the holotype (KU 308437, 335742��� 44) generally appear darker dorsally, owing to a combination of darker interstitial skin between the scales and fewer whitecolored scales. The holotype (KU 335744) and an additional specimen from Mindoro (KU 308437) were the largest intact specimens examined, and both exhibit a reduction in the size and vibrancy of dorsal banding, with the latter individual having bands that are only faintly discernable. Juvenile specimens from both Mindoro (KU 335742, 335743) and Semirara (KU 305151, 305161, 305163) exhibit much more distinct dorsal bands comprised of series of yellowish to white spots or oscelli. One juvenile from Mindoro (PNM 9799; formerly KU 308403) appears to be an intermediate, with dorsal bands discernable posteriorly on the trunk and becoming less defined anteriorly. One specimen from Semirara (KU 305161) exhibits a faded reticulate pattern interspersed among the more well-defined dorsal bands, owing to clusters of grayish spots comprised of lightly colored scales and interstitial space. Specimens from Semirara (KU 305151, 305161, 305163) and one from Mindoro (KU 335742) exhibit a more well-defined light-colored longitudinal stripe laterally in the nuchal region, which extends from the ventral margin of the tympanum dorso-posteriorly over the forelimb insertions, joining with the first dorsal trunk band to form a vague ���U��� shape when viewed from dorsal perspective. Dorsal nuchal coloration varies from uniform black in the holotype (KU 335774) and three paratypes (PNM 9799, KU 308437, 335772), the presence of 4���10 variable spots or blotches (KU 305151, 308161, 335743), to a more mottled and irregular combination of stipes and blotches posteriorly (KU 305163). One specimen (KU 308437) exhibits a reduction in white speckling on the dorsal surfaces of the limbs. Ventral coloration is moderately variable. The characteristic gular blotches or spots are present in all specimens, though the number and relative size of blotches is somewhat reduced in four specimens (KU 305161, 308403, 335742, 335743). Two specimens (KU 305151, 305163) exhibit much larger and more conspicuous gular spots. Ventral trunk coloration is less variable, with only three specimens (KU 305161, 308403, 308437) exhibiting reduced transverse dark bands, which generally fail to converge medially. Ventral coloration of the limbs is nearly identical among specimens, with only slight variation in the appearance of black color encroaching on the lateral margins of both fore- and hindlimbs, but generally appearing as partial transverse bands. The holotype (KU 335774) and three paratypes (KU 308437, 305151, 305161) exhibit a more irregular pattern on the margins of the ventral limb surfaces, with dark coloration appearing as an almost reticulate pattern. Ventral coloration of the tail is nearly uniform across specimens, with only minor variation in the extent of medial extension of dark bands. In all specimens, dark bands fail to coalesce anteriorly, but posteriorly to the terminus form a distinct alternating black and white pattern (see Figures 6 and 7 for additional information on color pattern variation). Coloration in life. Based on the only available image, KU 335742 (Fig. 8 B), and field notes of the authors. Dorsal ground coloration black, with transverse crossbands of irregularly shaped creamy white to light yellow ocelli from the posterior margin of the nuchal region through the hind limb insertion. Trunk with few, larger ocelli with darkened, ground-colored centers. Additionally, dorso-lateral margins of trunk with smaller, variably distributed light yellow spots or blotches. Lateral margins of head and nuchal region with conspicuous light yellow stripe, extending from the the posterior margin of the ocular, through the tympanum, and midway into the anterior portion of the nuchal region. An additional light yellow stripe extends dorsally and posteriorly from the ventrolateral margin of the nuchal region, and extends to the dorsal region above the forelimb insertion. Snout with distinct and conspicuous bands extending ventrally from dorsal surface. Dorsal limb surfaces colored similarly to body, but with ocelli irregularly arranged and significantly smaller, lacking any dark-colored center. Forelimbs with larger ocelli dorso-medially. Ventral coloration creamy white to light yellow, with scattered black pigmentation and with ocelli of ventrolateral surfaces irregularly arranged and significantly smaller, lacking any dark-colored center (Fig. 8 b). Ecology, Distribution, and Natural History.���As with all members of the Varanus salvator Complex, V. bangonorum should be considered a habitat generalist, often thriving in a range of habitats from primary forest to more urbanized areas. Like V. dalubhasa, no detailed studies have been undertaken on this taxon specifically, but the findings of Gaulke (in Luxmoore & Groombridge 1989, 1992) are likely relevant. This taxon is most likely to be found in mangrove and riparian habitats (as their common name might suggest). Additionally, most water monitors seek shelter in rock crevices or even trees, often near water sources. Not surprisingly, Varanus bangonorum has been found on the islands of Mindoro and Semirara (Figs. 1, 2), both of which are separated by deep water channels that likely have facilitated the isolation of this species and its evolution as a distinct lineage, even during inter-glacial cycles and lowered sea levels (Voris 2000; Brown & Diesmos 2009). Again, the findings of Gaulke (1992) are relevent as far as dietary preferences are concerned, and it can reasonably be assumed that the diet of Varanus bangonorum consists primarily of invertebrates and smaller vertebrates (fish, rodents, frogs, snakes). Additionally, as with any species of water monitor, this species is attracted to a number of carion items, with a particular affinity for fish (LJW pers. obs.)., Published as part of Welton, Luke J., Travers, Scott L., Siler, Cameron D. & Brown, Rafe M., 2014, Integrative taxonomy and phylogeny-based species delimitation of Philippine water monitor lizards (Varanus salvator Complex) with descriptions of two new cryptic species, pp. 201-227 in Zootaxa 3881 (3) on pages 216-221, DOI: 10.11646/zootaxa.3881.3.1, http://zenodo.org/record/226203, {"references":["Tweddell, C. E. (1970) The identity and distribution of the Mangyan tribes of Mindoro, Philippines. Anthropological Linguistics, 12 (6), 189 - 207.","Welton, L. J., Siler, C. D. Bennet, D., Diesmos, A. C., Duya, M. R., Dugay, R., Rico, E. L., van Weerd, M. & Brown, R. M. (2010) A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation. Biology Letters, 6, 654 - 658.","Luxmoore, R. & Groombridge, B. (1989) Asian monitor lizards: a review of distribution, status, exploitation and trade in four selected species. Report to the Cites Secretariat, World Conservation Monitoring Centre, Cambridge.","Voris, H. K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems, time durations. Journal of Biogeography, 27, 1153 - 1167.","Brown, R. M. & Diesmos, A. C. (2009) Philippines, Biology, In: Gillespie, R. & Clague, D. (Eds.), Encyclopedia of Islands. University of California Press, Berkeley, pp. 723 - 732.","Gaulke, M. (1992) Taxonomy and biology of Philippine water monitors (Varanus salvator). The Philippine Journal of Science 121 (4), 345 - 364."]}

Varanus dalubhasa sp. nov. Figs. 3, 5���8 Holotype. PNM 9796 (formerly University of Kansas Biodiversity Institute [KU] 306603; Field No. CWL 521), adult male, collected by Charles W. Linkem and CDS, 0 8 July 2006, 18 m above sea level (14.03202, 122.34143; WGS- 84), Barangay Madlangdungan, Municipality of Calauag, Quezon Province, Luzon Island, Philippines. Paratopotypes. KU 305155 (CDS Field No. 2202), adult male; PNM 9797 (formerly KU 306601; CWL Field No. 440), juvenile; KU 306602 (CWL Field No. 520), adult male. Paratypes. KU 308216 (CDS Field No. 2298), juvenile, collected 22 February 2007, Barangay Buenavista, Municipality of Bato, Catanduanes Province, Catanduanes Island, Philippines; KU 313880 (RMB Field No. 9910), adult male, collected 0 1 July 2008, Barangay Tulay Na Lupa, Municipality of Labo, Camarines Norte Province, Luzon Island; KU 326702 and 326703 (LJW Field No. 0 0 75, 0077), adult males, salvaged 3 August 2009, Polillo Island (exact locality information unknown). Other Material. See Specimens Examined Section. Etymology. The specific epithet, dalubhasa, is derived from the Tagolog word ��� dalubhasa ��� meaning a person who has authoritative and comprehensive knowledge of a particular area, or a skilled expert in a particular subject. We choose this term in honor of Vicente ���Enteng��� Yngente of Polillo Island, whose extensive knowledge of natural history and ecology of Philippine reptiles (particularly monitor lizards) has been instrumental to our research and conservation work. Suggested common name: Enteng���s Monitor Lizard. Diagnosis. Varanus dalubhasa can be distinguished from congeners by (1) small, dark speckling and variable transverse banding through the gular region (Fig. 3); (2) number of scales around the tail at 1 / 3 distance from the base; (3) number of gular scales; (4) number of dorsal scales in nuchal region; (5) total number of dorsal scales; (6) number of scales around the neck anterior to the gular fold; and (7) phylogenetic placement sister to V. nuchalis. Additionally, this distinct lineage is biogeographically circumscribed in the Bicol Peninsula faunal sub-region, a distinct geological component of greater Luzon Island which remained inuslar until 3 Ma (Hall 2002). Comparisons. The new species in phenotypically nearly indistinguishable from Varanus marmoratus, but can generally be diagnosed by the presence of small, dark speckling in the gular region and faint anterior transverse bands (versus speckling and distinct anterior transverse bands; Fig. 3), and its allopatric distribution in the Bicol faunal sub-region (versus the remaining portions of Luzon and Lubang islands, and the Batanes and Babuyan island groups). Additionally, although V. dalubhasa is phenotypically similar to V. marmoratus, available data suggest that it is not most closely related to this species (Fig. 2). Varanus dalubhasa can be conveniently distinguished from its allopatric sister taxon, V. nuchalis (distributed in the West Visayan islands of Negros, Panay, Guimaras, Masbate, and the Romblon Province islands of Sibuyan, Tablas, and Romblon; Fig. 1, 2), by the presence of more scales around the tail at 1 / 3 distance from the base (mean = 53 �� 4, versus 46 �� 3), fewer gular scales (mean = 80 �� 3, versus 73 �� 3), and more dorsal scales from the tympanum to the gular fold (mean = 27 �� 2, versus 23 �� 2). Of the remaining, geographically proximate species of the V. salvator Complex, this new species can be distinguished from V. palawanensis by having fewer dorsal scales from the tympanum to the gular fold (mean = 27 �� 2, versus 32 �� 3), fewer dorsal scales from the gular fold to the hind limb insertion (mean = 84 �� 5, versus 96 �� 3), fewer total dorsals scales longitudinally (mean = 111 �� 5, versus 128 �� 3), and fewer scales around the neck anterior to the gular fold (mean = 78 �� 3, versus 87 �� 4). Additionally, V. dalubhasa can be distinguished from the allopatric Mindoro lineage by having variable dark speckling and transverse bands in the gular region (versus distinct spotting; Fig. 3). Lastly, V. dalubhasa is comprised of entirely distinct haplotypes or haplotype networks, relative to all other members of the V. salvator Complex (see Welton et al., 2010 a). We have constrained our morphological analyses to the geographically most proximate taxa due to previous studies (Koch et al. 2007, 2010 b) demonstrating their distinctiveness relative to the remaining diversity withing the V. salvator Complex. Description of holotype. An adult male (Fig. 5), hemipenes everted; SVL 505.0 mm; tail 657 mm; head relatively slender, length 89.1 mm, width 51.0 mm, and depth at eye 32.0 mm; snout length 47.6 mm, 53.4% head length; snout rounded anteriorly; narial openings 5.4 mm, ovular, posteriorly elevated, encircled by 9 L/ 9 R small polygonal scales; snout elevated sharply just anterior to narial region, then continuing gradually to ocular region; narial region not distinctly elevated above horizon of snout; cranial table squarish, only slightly wider than long. Head scales ovular to polygonal (Fig. 5), heterogeneous in size with the smallest occurring dorso-laterally between ocular opening and tympanum, and largest ocurring dorso-medially from tip of snout to conspicuous parietal; supraoculars 6 L/ 6 R, subrectangular and elongated transversely; supralabials 59, slightly larger than lateral head scales; infralabials 61, smaller than both supralabials and nearest rows of gular scales; both supra- and infralabials increasing in size anteriorly; nuchals large, circular, increasing in size to forelimb insertion along both dorsal and lateral surfaces; lateral nuchals less than half the size of dorsals; scales immediately posterior to cranial table quite small, granular; sutures between scales slightly larger in nuchal region than on head; nuchals 31, in semi-regular transverse rows from posterior margin of tympanum to forelimb insertion; dorsal trunk scales arranged in 88 semi-regular transverse rows from fore- to hindlimb insertions, ovular, slightly smaller than nuchals, gradually decreasing in size to hindlimb insertion as well as laterally, with smallest scales occuring laterally adjecent to limb insertions; midbody scales 138; axilla���groin distance 245 mm; dorsal limb scales ovular, decreasing in size distally to digits; fore- and hindlimbs 92 and 107 mm, respectively, 18.2% and 21.2% snout���vent length, 32.6 and 53.9 mm diameter at insertions; limb scales present in semiregular rows; scales of manus and pes smaller and more irregular in shape, ranging from circular to nearly rectangular dorsally; supradigitals rectangular dorsally, ovular laterally, with terminal scale nearly twice as large as others; digits terminate in long, slender recurved claws; caudals ovular and granular at insertion, becoming rectangular posteriorly; dorsal keel composed of paired, posteriorly elevated scales commenses 65 mm posterior to hindlimb insertion, and continues to tail terminus with paired scales gradually decreasing in size; caudal scales moderately keeled, in regular transverse and longitudinal rows laterally, decreasing in size posteriorly. Ventrals less variable than dorsals; gular scales generally rectangular anteriorly, becoming more ovular towards gular fold; smallest scales medially in anterior third of gular region, with largest scales just anterior to gular fold but followed posteriorly by three transverse rows of smaller, more granular gulars; scales around the neck anterior to the gular fold 78, mid-gular 87; gular scales in 85 irregular rows from tip of snout to gular fold; ventrals of fore- and hindlimbs circular to polygonal, with those of forelimbs slightly smaller than those of hindlimbs; precloacals homogenous in size and shape with those of hindlimbs; ventral trunk scales from gular fold to hindlimb insertion 85, ovular anteriorly, becoming rectangular posteriorly, and present in semi-regular rows; scales decreasing in size laterally and at hindlimb insertions; ventral caudal scales rectangular, smallest just posterior to cloacal opening; scale surface slightly rounded anteriorly, becoming sharply keeled and more longitudinally elongate posteriorly. Right and left hemipenes partially everted; maximum length 56.2 mm, with proximal cylindrical section extending 26.2 mm; bifurcation resulting in primary cylindrical apical portion with external flounces, and more diminutive portion resembling an ���ear��� or reduced fold; flounces present in irregular rows, totalling 15 anterior to sulcus spermaticus, 6 posteriorly; lacking notable external appearance of ossified hemibaculum. Measurements of holotype (in mm). SVL 505; tail length 657; head length 89.1, width 51.0, depth at eye 32.0; eye���naris distance 33.7; naris���snout distance 14.9; rictus���rictus scales 60; scales around tail base 103; scales around tail 1 / 3 distance form base 59; mid-body scales 138; gular scales 85; ventrals from gular fold to hind limb insertion 85; total ventral scales 170; dorsals from tympanum to gular fold 31; dorsals from gular fold to hind limb insertion 88; total dorsals 119; supralabials 59; scales around the neck anterior to gular fold 78; and enlarged supraoculars 6 L/ 6 R. Coloration of holotype in preservative. Dorsal surfaces of head, body, and limbs black; aggregation of yellow-gold scales present dorsoanteriorly to narial openings, along canthal ridge just anterior to oculars, and irregularly dispersed throughout cranial table; trunk with seven transverse rows comprised of yellow-gold, indistinct ocelli; tail with nine, slightly more distinct yellow-gold bands, becoming more diffuse or speckled laterally; forelimbs with irregularly distributed yellow-gold scales, density increasing distally and along lateral surfaces; hindlimbs with irregular agreggations of ocelli consisting of only two or three yellow-gold scales (Fig. 5). Lateral coloration of head consisting of partial, near vertical yellow-gold bands at the tip of snout, posterior margin of narial opening, and two between the narial opening and the ocular; posterior band with ventral widening oriented more posteriorly; post-orbitally, a yellow-gold stipe extends to tympanum, and a yellow-gold spot consisting of four or five scales appears ventral to the anterior margin of the stripe; lateral coloration of the nuchal region consists of 2 L/ 3 R faint spots directly posterior to the tympanum; ventral margins of nuchal region with yellow-gold variable demarcation, extending more dorsally along the posterior margin and appearing to wrap around the dorsal margin of the fore-limb insertions; trunk with variable yellow-gold spotting directly posterior to forelimbs, fading to solid black posteriorly (Fig. 5). Ventral coloration predominately yellow-gold throughout; gular region with irregular spots, faint anteriorly but becoming prominent and dark along lateral margins just anterior to gular fold; scapular region with two irregular black bands; trunk with irregular black bands and spots anteriorly, becoming more broken medially and posteriorly, appearing as laterally elongated yellow-gold ovals; forelimbs with faint, irregular dark mottling; hindlimbs with near uniform venter medially, with dark mottling along lateral margins; manus and pes dark yellow-gray; venter of tail appearing anteriorly much like that of the trunk, with yellow-gold laterally elongated ovals; posterior portions of tail with more regular alternation of dark and yellow-gold bands (Fig. 5). Variation. Our small type series exhibits some color variation, but specimens are generally phenotypically similar. The holotype (KU 306603) and three paratypes (KU 305155, 306602, and 308126) have lighter interstitial coloration between the dorsal scales, yielding an overall lighter appearance than other specimens. These three specimens also exhibit much more diffuse yellow gold coloration on limbs and tail, while two specimens (KU 313880, 326702) exhibit much more distinct yellow-gold spots or ocelli throughout. Additionally, the yellow-gold coloration in the lateral portions of the nuchal region, among all specimens, and on either side of a single specimen, are present as either a single longitudinal stripe extending posteriorly from the posterior margin of the tympanum or as a series of two or three longitudinally elongated spots or blotches. Two specimens (KU 305155, 306602) exhibit two relatively well-defined yellow crossbands on the snout, with the former���s being even more pronounced than the latter, and with this specimen (KU 305155) also exhibiting a much larger proportion of yellow coloration in the supraocular and temporal regions. In contrast, two specimens (KU 308216, 313880) exhibit nearly uniform, dark dorsal head coloration, with the former being completely devoid of yellow-gold accents with the exception of the pineal scale. Ventral coloration is only slightly more variable than that of the dorsum. All specimens exhibit variable speckling in the gular region, gernerally consisting of 1���4 black scales. In the holotype (KU 306603), this speckling is much more reduced. In one specimen (KU 305155) anterior speckling in the gular region gives way to 2 broken transverse bands, and somewhat reduced speckling through the remainder of the gular region. Three specimens (KU 306602, 308216, and 326702) exhibit four, more well-defined dark transverse bands in the anterior portion of the gular regions, with speckling throughout the remainder of the gular region being relatively dense. A single specimen (KU 313880) exhibits nearly complete dark transverse bands throughout the gular region, with less prominent bands present anteriorly. Ventral trunk coloration is nearly uniform among specimens, with the only notable variation present in two individuals (KU 305155, 326702), the first of which exhibits reduced dark coloration extending medially onto the abdomen, and the second of which with darker, more distinct coloration extending medially onto the abdomen. Ventral tail coloration lacks any notable variation among specimens (see Figures 6 and 7 for further information on color pattern variation). Coloration in life. Based on the only available image, PNM 9797 (formerly KU 306601; Fig. 8 A), and field notes of the authors. Dorsal ground coloration black to dark gray, with consipcuous transverse cross bands of yellow-gold ocelli from the posterior margin of the nuchal region through the anterior third of the tail. Trunk ocelli larger with a darkened, ground-colored center. Additionally, dorsal trunk with faded reticulate pattern, owing to light brown interstitial coloration. Lateral margins of head and nuchal region with conspicuous yellow-gold stripe, extending from the the posterior margin of the ocular, through the tympanum, and midway into the nuchal region. Yellow-gold ocelli brightest at lateral margins of trunk and adjacent to hindlimb and tail insertions. Dorsal limb surfaces colored similarly to body, but with ocelli irregularly arranged and significantly smaller, lacking any dark-colored center. Dorsal ground color of limbs bright yellow, with irregular dark pigmentation. Ventral coloration bright yellow-gold, with scattered black pigmentation and with ocelli of ventro-lateral surfaces irregularly arranged and significantly smaller, lacking any dark-colored center (Fig. 8 A). Ecology, Distribution, and Natural History. As with all members of the Varanus salvator Complex, V. dalubhasa should be considered a habitat generalist, often thriving in a range of habitats from primary forest to more urbanized areas. While no detailed studies have been undertaken on this taxon specifically, the findings of Gaulke (in Luxmoore & Groombridge 1989, 1992) are likely relevant. This taxon is most likely to be found in mangrove and riparian habitats (as their common name might suggest). Additionally, most water monitors seek out shelters in the form of rock crevices or even trees, most often near water sources. In more urban areas, human habitation may be used in lieu of more ���natural��� shelters (Gaulke 1992). Not surprisingly, Varanus dalubhasa has been found throughout the Bicol Peninsula and faunal sub-region, including genetically confirmed specimen identifications from Polillo and Catanduanes islands (Figs. 1, 2). This is an area with significant agricultural (rice fields) and coastal environments, and is additionally subdivided by a number of moderately sized river drainages. The northern extent of distribution for this taxon, and potential contact zone with V. marmoratus requires further study. Although no dietary preferences have been observed for this taxon, one can reasonably assume that its diet consists primarily of invertebrates and smaller vertebrates (fish, rodents, frogs, snakes). Additionally, this species is attracted to a number of carion items, with a particular affinity for fish (LJW pers. obs.)., Published as part of Welton, Luke J., Travers, Scott L., Siler, Cameron D. & Brown, Rafe M., 2014, Integrative taxonomy and phylogeny-based species delimitation of Philippine water monitor lizards (Varanus salvator Complex) with descriptions of two new cryptic species, pp. 201-227 in Zootaxa 3881 (3) on pages 212-215, DOI: 10.11646/zootaxa.3881.3.1, http://zenodo.org/record/226203, {"references":["Hall, R. (2002) Cenozoic geological and plate tectonic evolution of SE Asia and the SW Pacific: computer-based reconstructions, model and animations. Journal of Asian Earth Sciences, 20, 353 - 431.","Welton, L. J., Siler, C. D. Bennet, D., Diesmos, A. C., Duya, M. R., Dugay, R., Rico, E. L., van Weerd, M. & Brown, R. M. (2010) A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation. Biology Letters, 6, 654 - 658.","Luxmoore, R. & Groombridge, B. (1989) Asian monitor lizards: a review of distribution, status, exploitation and trade in four selected species. Report to the Cites Secretariat, World Conservation Monitoring Centre, Cambridge.","Gaulke, M. (1992) Taxonomy and biology of Philippine water monitors (Varanus salvator). The Philippine Journal of Science 121 (4), 345 - 364."]}

Cyrtodactylus jambangan sp. nov. Figs. 3–5 Holotype. PNM 9593 (formerly KU 314810, RMB Field No. 10,005), adult male, collected on the trunk of a sapling (20:00– 23:00) in Pasonanca Natural Park, Barangay Baluno, Zamboanga City Province, Mindanao Island, Philippines (N: 07º 1.0554, E: 122 º 1.731, WGS- 84; 758 m above sea level), on 12 July, 2008 by L. Welton, R. Brown, C. Siler, J. Fernandez, M. and V. Yngente, and J. Phenix. Paratopotypes. Four males (KU 314813, 314815, 314825, and 314828), fourteen females (KU 314808, 314811, 314816 – 18, 314820, 3134822 – 24, 314826–27, and 314829 – 31), and five juveniles (KU 314809, 314812, 314814, 314819, and 314821) collected on the trunks of trees (18:00– 23:00), 12–18 July 2008, other data identical to holotype. Paratypes. Four males (KU 314793 – 94 and 314796 – 97), two females (KU 314795 and 314798), and three juveniles (KU 314792, and 314833 – 34), collected 7 July 2008, in Pasonanca Natural Park, Tumaga River (N: 6 º 35.174, E: 122 º 2.426, 94m, WGS- 84), by L. Welton, R. Brown, C. Siler, J. Fernandez, M. and V. Yngente, and J. Phenix; one male (KU 314835), one female (KU 314806), and one juvenile (KU 314799), collected 8 July 2008, in Pasonanca Natural Park, Tumaga River (N: 6 º 58.624, E: 122 º 3.043, 104m, WGS- 84), by the same collectors; two males (KU 314807 and 314836) collected 9–11 July 2008, in Pasonanca Natural Park, Tumaga River (N: 6 º 59.534, E: 122 º 3.626, 94m, WGS- 84), by the same collectors; two males (KU 314803 and 314805), one female (KU 314804), and three juveniles (KU 314800 –02), collected 9 July 2008, Pasonanca Natural Park, Tumaga River (N: 6 º 58.624, E: 122 º 4.043, 104m, WGS- 84), by the same collectors; two males (KU 314781 and 314784), one female (KU 314780), and four juveniles (KU 314778 – 79 and 314782 – 83), collected 20–22 April 2008, in Pasonanca Natural Park, Barangay Pasonanca, Sitio Canucutan (N: 6 º 59.534, E: 122 º 3.626, 104m, WGS- 84), by R. Brown and A. Diesmos; one male (KU 319654), one female (KU 319655), and one juvenile (KU 319652), collected 3–5 April 2009, in Pasonanca Natural Park, Barangay La Paz, Sitio Nancy (N: 7 º 5.099, E: 122 º 1.620, 1130m, WGS- 84), by R. Brown, C. Infante, and A. Diesmos; two males (KU 319656 and 319657), collected 6–9 April 2009, in Pasonanca Natural Park, Barangay Tulosa, Sitio Santa Clara, Cabo Negros outpost (N: 7 º 6.479, E: 122 º 7.139, 620m, WGS- 84), by the same collectors; three males (KU 319660 – 62), and one female (KU 319658), collected 11 April 2009, in Pasonanca Natural Park, Tumaga River (N: 6 º 35.174, E: 122 º 2.426, 94m, WGS- 84), by the same collectors; twelve males (CAS 60195, 60197, 60199, 60203 – 205, 60208, 60212, 60217 – 18, 60221, and 60453), 15 females (CAS 60196, 60200 – 201, 60207, 60209 – 10, 60213–16, 60220, 60454 – 55, 60458, and 60460), and six juveniles (CAS 60222 – 24, 60456–57, and 60459) collected at Abung-Abung and Port Holland on Basilan Island, 5–25 October, 1921, by E. H. Taylor; two males (CAS 60622 – 23), three females (CAS 60619 – 21), and one juvenile (CAS 60624) collected on New Govenen Island, 5–25 October, 1920, by E. H. Taylor; one male (CAS 60669), four females (CAS 60670 – 72 and 60887), and two juveniles (CAS 60886 and 60888) collected on Bud Daho Mt. on Jolo Island, 25 October– 17 November, 1920, by E. H. Taylor; two males (CAS 62020 and 62022), three females (CAS 62017, 62019, and 62021), and one juvenile (CAS 62018) collected in Zamboanga del Sur Province, 23 September– 6 October, 1920, by E. H. Taylor. Diagnosis. The new species differs from most of Asia’s 115 Cyrtodactylus species by the same suite of character states that has facilitated the recognition and distinction of C. annulatus: (1) small body size; (2) light gray body, with dark transverse bands; (3) presence of a weak precloacal groove; (4) absence of enlarged femoral scales and pores; (5) moderately spinose tuberculation; (6) presence of forelimb tuberculation; (7) and absence of enlarged, median subcaudal scales (Taylor 1922; Brown & Alcala 1978; Manthey & Grossman 1997; Grismer 2005; Welton et al. 2009). Because the new species differs from non-Philippine congeners by characters that have distinguished C. annulatus (and have never been challenged; Taylor 1922 a; Brown & Alcala 1978), we do not provide exhaustive comparisons to all those species (see Welton et al. 2009). Instead we focus on the comparisons relevant for the recognition of the new species: the closely related taxa within the C. annulatus complex and other Cyrtodactylus endemic to the Philippines. C yrtodactylus jambangan differs from all other species of Philippine Cyrtodactylus (i.e., C. agusanensis, C. philippinicus, C. redimiculus, and C. tautbatorum; Table 2) by the following combination of characters: (1) small body size (2) dorsum darkly marbled lavender and brown or brown with indistinct wavy lavender blotches; (3) presence of bright yellow superciliaries, canthal stripe, and dorsal tubercles; (4) low dorsal scale counts (as measured by midbody dorsal and paravertebral scales; Table 2); (5) moderate numbers of midbody dorsal tubercle rows (Table 2); (6) absence of femoral pores in both sexes; (7) presence of precloacal pores in males and similarly enlarged, dimpled, scales in females, arranged in an inverted “V”-shaped configuration; (8) moderately depressed precloacal groove; (9) scales anterior to precloacal region undifferentiated; (10) lamellae under Finger III 17–22; (11) lamellae under Toe IV 20–24 (12) supralabials to beneath eye 8–11; (13) infralabials to beneath the eye 6–8; (14) absence of stripes connecting the lateral margins of transverse bands on trunk; and (15) midbody ventral scales 48–63. Comparisons with additional Southeast Asian species are provided in Table 3. The critical comparison for the recognition of the new species is the diagnosis of the Zamboanga and Sulu populations from true C. annulatus (type locality from the Municipality of Bunawan, Agusan Del Sur Province, Mindanao Island), and newly designated C. annulatus neotype locality (Welton et al. 2009) of Mt. Hilonghilong, Diwata Mountains, Agusan Del Norte Province, Mindanao Island. Although morphologically similar to C. annulatus from throughout its range, C. jambangan has bright yellow canthal stripes, superciliaries, dorsal tubercles, and moderate-sized dorsolateral body tubercles, fewer precloacal pores, undifferentiated precloacals, and strongly spinose (versus moderate, domed) dorsal tuberculation. The new species is separated from C. annulatus by 7.9–9.5% (mean = 8.6%) uncorrected sequence divergence (Table 4) in the ND 2 protein-coding mitochondrial gene region (see Table 3 for complete univariate morphometric data for C. annulatus and C. jambangan). The new species differs from all remaining Philippine species by a variety of morphological and color pattern characters (Table 2; Brown and Alcala 1978). Description of holotype. Adult male, snout–vent length 67.7 mm; head moderately long, distinct from neck, 30.3% SVL; head width 64.7% and height 41.7% head length; head triangular in dorsal aspect; lores concave; snout elongate, 41.7% head length, anterior tip rounded; eye diameter 20.6% head length, 76.3% eye–ear distance; auricular opening ovoid, longest axis 9.5% head length. Dorsal head scalation heterogenous, scales small and granular, with tubercle density sparse medially, tubercles increasing in size and density posteriorly and laterally; superciliaries increasing in size (laterally elongated) anteriorly, with largest scales at anterodorsal margin of orbit; rostral taller than wide, divided dorsally by inverted “Y” shaped crease; rostral bordered by large, anterior internasal and smaller, paired posterior internasals, supranasals, and first supralabials; internarial distance 2.2 mm; nostrils bordered by supranasals, rostral, 2 postnasals, and first supralabials; supralabials rectangular, decreasing in size posteriorly (left/right): 9 / 10 to midpoint of eye, 12 / 12 total; infralabials 7 / 7 to midpoint of eye, 11 / 10 total; supralabials bordered dorsally by secondary, slightly differentiated row of scales, extending to anterior margin of orbit; infralabials bordered ventrally by similar row, extending to anterior margin of orbit. Ventral head scalation heterogenous; mental bordered by first infralabials, and paired postmentals, with triangular projection posteriorly; 7 differentiated gular scales posterior to postmentals; first infralabial in contact with postmental and single differentiated gular; remaining gulars heterogenous, small and granular, increasing in size in nuchal region. Body elongate, axilla–groin distance 44.5% snout–vent length, lacking distinct ventrolateral folds; dorsal scalation granular, heterogenous, with semi-regular rows of rounded, posteriorly-oriented tubercles, increasing in protuberance (from convex to conical) and size posterior to hind limb insertions; tubercles in 17 longitudinal midbody and 27 paravertebral rows; paravertebral scales 164; transverse midbody dorsal and ventral scales 91 and 54 respectively, between lateral tubercle rows; ventral scales imbricate, slightly larger than dorsals and increasing in size medially; differentiated precloacals 4, arranged in an inverted “V” configuration pierced with enlarged pores, surrounding shallow and longitudinally short precloacal groove; two patches of enlarged scales anterolateral to precloacals (14 left, 13 right); two rows of slightly enlarged scales anterior to precloacals; three slightly enlarged scales posterior to precloacals. Forelimbs slender, forearm and upper arm 14.3% and 12.4% snout–vent length respectively; scalation on dorsal surfaces of forelimbs heterogenous, scales larger than ventrals; tubercles absent on upper arm, sparse on forearm, less spinose than dorsal trunk tubercles; ventral scalation homogenous, lacking tubercles; fingers well developed; Fingers I and III 37.4% and 62.2% forearm length, respectively; lamellae enlarged and slightly raised, lamellae proximal to inflection larger than those distal to inflection, slight increase in size from inflection to claw; finger number followed by subdigital lamellae (in parentheses): I (11), II (14), III (20), IV (18), V (15); all fingers clawed; claws well developed, sheathed by single dorsal and ventral scale. Hind limbs relatively robust, limb diameter at insertion twice that of forelimb; femur long, 16.8% snout– vent length; dorsal hindlimb scalation heterogenous, scales increasing in size distally, with tubercles regularly distributed, increasing in size and becoming more spinose distally; ventral scalation heterogenous, increasing in size distally, lacking tubercles; right limb with two conspicuous patches dorsally, lacking scales or tubercles (possibly resulting from injury); enlarged femoral scales (and pores) absent; toes well developed, relatively longer than those on manus; Toes I and IV 31.2% and 85.4% tibia length, respectively; lamellae enlarged and slightly raised, lamellae proximal to inflection larger than those distal to inflection, slight increase in size from inflection to claw; toe number followed by subdigital lamellae (in parentheses): I (12), II (16), III (20), IV (23), V (22); all toes clawed; claws well developed, sheathed by single dorsal and ventral scale. Tail original, length 78.2 mm, width and height 5.6 and 4.8 mm at base respectively, tapering abruptly posterior to hemipenal bulge, then gradually to terminus; 7 annuli discernable dorsally through lateral margins, posterior to which whorls cannot be distinguished, owing to absence of differentiated scales associated with annuli; tubercles smaller and less spinose than those of trunk; enlarged medial subcaudal scales absent; 1 enlarged post-cloacal tubercle present on each side of cloaca; anterior margin of hemipenal bulge with paired, laterally expanded, postcloacal glandular openings. Measurements of holotype (in mm). Snout–vent length 67.7; lamellae under Finger III 20; lamellae under Toe IV 23; Finger I length 3.6; Finger III length 6.0; Toe I length 3.3; Toe IV length 9.0; eye-narial distance 6.5; eye–ear distance 5.6; interorbital distance 2.6; snout length 8.6; internarial distance 2.2; eye diameter 4.2; upper arm length 8.4; upper arm length 9.7; femur length 11.4; tibia length 10.5; hand length 9.1; foot length 10.9; axilla–groin distance 10.1; tail length 84.6. Coloration of holotype in preservative. Dorsal ground coloration (of head, neck, body, limbs, and tail) dark brown, with irregularly shaped dark gray to lavender transverse dorsal bands, varying from irregular dark gray blotches anteriorly, to a single medial blotch followed posteriorly by a “V-shaped” marking in the nuchal region, to three conspicuous, “butterfly-shaped” patterns interspersed with three irregularly shaped transverse blotches throughout the trunk, and becoming less-defined laterally; supralabials with four conspicuous light tan to white blotches (left and right) irregularly distributed between the second and twelfth scales; light gray band bordered by dark brown extends from posterior margin of eye to auricular opening; trunk tubercles primarily light tan to cream; limbs mottled, with dark gray and dark brown, lacking distinct bands; digits with light tan to cream blotches at joints; tail more conspicuously banded than trunk, with dark brown ground coloration fading to white at the terminus of tail, overlain with 10 dark gray to black bands. Ventral portions of head, trunk, and limbs tan, darkening at margins of ventrolateral tubercle row; hands and feet tan, fingers and toes slightly darker; subcaudal coloration cream with increased dark speckling through anterior third; seven discernable white bands posteriorly. Color Variation. Our sample of 36 males, 52 females, and 26 juveniles exhibits minimal color variation. Four adult males (KU 314794, 314796, 314813, and 314825) and four adult females (KU 314804, 314823– 24, and 314831) are darker, with dorsal bands through the axilla-groin region of higher contrast to the lighter ground color. One adult male (KU 314815) and two adult females (KU 314822, and 314827) are lighter, with distinct dorsal banding, and light brown to light gray ground color overlaid by medium to dark gray band. One adult male (KU 314805) and one adult female (KU 314795) lack a defined “V-shaped” blotch spanning the posterior dorsum of the head and nuchal region. Three adult males (KU 314794, 314805, and 314835) lack the conspicuous blotch medially in the nuchal region. All other specimens have color and patterning consistent with the holotype. Additionally, nine adult males (KU 314781, 314784– 85, 314789, 314791, 314797, 314803, 314812, and 314836), 19 adult females (KU 314778 – 80, 314787–88, 314790, 314792, 314798, 314808 –09, 314816 – 19, 314820–21, 314829, 314832, and 314834), and 10 juveniles (KU 314782 – 83, 314883, 314799 – 802, 314807, 314786, and 314814) show similar trends in variation, with one adult male (KU 314797), two adult females (KU 314816 and 314832), and two juveniles (KU 314814 and 314833) with a darker overall coloration and more highly defined dorsal banding. Four adult males (KU 314781, 314784, 314789, and 314803) and 13 adult females (KU 314734, 314779– 80, 314790, 314792, 314798, 314809, 314817 – 21, and 314829) have a light gray ground color with dark gray, well-defined dorsal banding. Regenerated tails have highly variable dark speckling overlaying a medium gray ground color, to a completely dark brown dorsal coloration. Generally, a greater level of definition in dorsal banding is present in female and juveniles; these specimens have greater contrast in dorsal banding; a few individuals nearly lack dorsal bands all together (KU 314799, 314801, and 314802). Ventral coloration is less variable except on the tail. Original tails are similarly colored with speckling increasing from the anterior margin of the tail to the terminus, with the tail tip being nearly black. Dorsal, light-colored bands extend through venter, becoming lighter in color and more defined towards terminus of tail. One adult male (KU 314825) has speckling at the base and the tip of the tail, a cream ground color throughout, and lacks dark patterning in between. All others exhibit increasing amounts of dark speckling over a light to medium gray ground color from the anterior margin of the tail to the terminus. Morphological variation. Summaries of univariate morphological variation in the series are presented in Table 3. Our series shows minimal morphological variation between males and females. However, females are generally larger and more robust (the largest specimen, KU 314831, is female with SVL = 81.5). Color in life. Cyrtodactylus jambangan has light yellow to gold dorsal tubercles through the axilla–groin region, a yellow canthal stripe, bright yellow superciliaries, and a variable dorsal banding pattern. All three characters are in contrast to observations of true C. annulatus, which has regular, distinct, and brightly contrasting dorsal bands through the axilla-groin region, more pattern variation, and also lacks bright yellow canthal stripe and superciliaries (see Fig. 5). Ecology and natural history. The new species is found at low and mid-elevations in riparian habitats (gallery forests) along streams. We encountered C. jambangan specimens on trunks of trees, rocks, and overhanging exposed root masses on the banks of rivers and streams. Distribution. The distribution of the new species includes Pasonanca Natural Park, Zamboanga Peninsula, extreme western Mindanao Island, and adjacent Sulu Archipelago islands to the southwest (Fig. 1). Other localities in the Sulu Archipelago where Taylor (1922 a) observed this species include Basilan, Great Santa Cruz, Teipono, Tamuk, Cancuman, Dipolod, Bitinan, Jolo, Tulian, Tawitawi, Papahag, Bongao, Buban Islands of the Tapiantana and Taipan Island Groups. Etymology. The specific epithet is derived from the term Jambangan —the ancient name for the Zamboanga City area. Sulu and Zamboanga folklore suggest that the name was bestowed upon the Zamboanga City area by the immigrant Subanons (“People of the River”) who arrived in western Mindanao and the Sulu archipelago in approximately 1200 A.D., after traveling by boat through the Sulu Archipelago, from what is now Indonesia. Jambangan means “The Land of Flowers,” a name presumably used in reference to the natural beauty of the area surrounding Zamboanga City.